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The Biogeography and Ecology of the Secondary Marine Arthropods of Southern Africa \

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The Biogeography and Ecology of the Secondary Marine Arthropods of Southern Africa \ The biogeography and ecology of the secondary marine arthropods of southern Africa \ . by ~erban Proche~ Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Degree in the School of Life and Environmental Sciences Faculty of Science and Engineering University of Durban-Westville Promoter: Dr. David J. Marshall November 2001 DECLARATION The Registrar (Academic) UNIVERSITY OF DURBAN-WESTVILLE Dear Sir I, Mihai ~erban Proche§ REG. NO.: 9904878 hereby declare that the thesis entitled The biogeography and ecology of the secondary marine arthropods of southern Africa is the result of my own investigation and research and that it has not been submitted in part or full for any other degree or to any other University. S.tl"h"iA. ~('oc~ c· ----- ~ ------------------------ ~ 15 November 2001 Signature Date 11 The biogeography and ecology of the secondary marine arthropods of southern Africa ~erban Proche§ Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy degree in the School of Life and Environmental Sciences, Faculty of Science and Engineering, University of Durban-Westville, November 200l. Promoter: Dr. David J. Marshall. Abstract Because of their recent terrestrial ancestry, secondary marine organisms usually differ from primary marine organisms in life history and physiological traits. Intuitively, the traits of secondary marine organisms constrain distribution, thus making these organisms interesting subjects for comparative investigation on ecological and biogeographical theory. A primary objective of the studies presented here was to improve our current knowledge and understanding of the generally poorly known secondary marine arthropods (e.g. mites and insects). An additional objective was to outline relationships between ancestry, ecology, and biogeography of small-bodied, benthic marine arthropods. In establishing a context for the global biogeographical study, the distribution patterns of secondary marine plants and animals were determined by means of a literature survey. These organisms, including mangrove trees and marine tetrapods, form three distinct groupings which relate to northern, tropical and southern latitudinal bands, and, exhibit bimodal species richness in each hemisphere. The exact same patterns were resolved for non­ halacarid marine mites and are thought to have arisen from the effects of elevated tropical speciation and high latitude glaciation. Whereas the typically marine Halacaridae show a high degree of radiation in sub-tidal habitats, global distribution patterns and species to genus ratios indicate that the intertidally-restricted Ameronothroidea (Oribatida) and Hyadesiidae (Astigmata), have had long marine associations relative to other non-ha1carid mites (Mestostigmata and Prostigmata). The mari~e mite fauna of southern Africa clusters into three geographical provinces, and the species richness of rocky shore mites peaks in the southern, warm temperate province. In being consistent with the trend for the region'S marine fauna in III general, the mite biogeography highlights the generality of this faunistic trend, with respect to taxonomic resolution and taxonomic diversity. Ecological studies focussing on mangrove pneumatophores show that they support a characteristic suite of arthropods (mites, copepods, tanaids, insect larvae), which differs from that of the benthic sediment. Pneumatophore assemblages comprise similar numbers of primary and secondary marine species, although the former group is more abundant by one order of magnitude. Pneumatophore assemblage composition varies between mangrove forests, predominantly in relation to salinity variation. Within mangrove forests differences arise through differential wetting frequency and variable sunlight intensity. Desiccation­ limited algal growth and sediment deposition determine the vertical zonation of arthropods along the pneumatophore length, with secondary marine species typically occurring at lower elevations, and primary marine species at higher elevations. The levels of similarity in the composition of arthropod assemblages decreases with spatial scale, the largest differences occurring between estuarine systems. Despite their recent terrestrial ongms, secondary marine arthropods show a high degree of integration into marine ecosystems, and should be consistently considered in marine ecological and biogeographical studies. Key Terms: Acari, ecological transitions, intertidal, mangroves, mites, southern Africa IV Acknowledgements Lots of thanks to my promoter, David 'Mick' Marshall, who took the chance of offering a bursary to an unknown student, coming from faraway Romania. If I'll ever be anywhere in the scientific world, I will owe it to him. Thanks to the NRF and UDW for funding me, and to the MCM for granting me collection permits. My gratitude to Profs. Carolyn Baker and Ahmed Thandar for their easy-going approach to administrative matters, and to the technical staff for taking me around on collecting trips. Thanks to Angelo Lambiris and Renzo Perissinotto for amazing exchanges of scientific, and not only scientific, ideas. Thanks in their respective languages to all my excellent friends here in Durbs. They made these two years and a half, more than a PhD, a real social experience. I specifically want to name here Santosh Bachoo, Dennis Ball, Amar Bholonath, Ketelo Hlehlisi, Israel Kibirige, Deshi Moodley, June Munnisunker, Mags Natasen-Moodley, Phindi Nonyane, Christian Nozais, Ashvena and Ashvita Ramcharan, Syd Ramdhani, Vanessa Singh, and Kaajial U grasen. I thought it appropriate to list here (in chronological order) the five people whose contribution was essential in my choice of natural science as a career. Mr. ~tefan Proche~ - my father, that is. He fashioned my first insect tray when I was four. He had reasons to get seriously worried later on, when another seven were required to accommodate my growing collections. Mr. Mihai Filip - my best friend from the gymnasium years, and my opponent in a fierce competition for 'who gathers more bugs' . Mr. Alexandru Iftime - my twin-brother-in-fate, who introduced me to the mysterious world of scientific names and encyclopedic knowledge. Mrs. Rodica Serafim - 'mom' of all Romanian beetles. She taught me the patience to go through endless pages of scientific literature, and grew me up as a real coleopterist. If I turned out otherwise, it's all my fault. Prof. Irina Teodorescu - my Honours and MSc supervisor. A lady of an almost extinct kind, she went through great pains explaining me that in science, to exist, one must publish. This thesis is dedicated to my parents, Ileana and ~tefan Proche~. Thanking them here for bringing me up would be beside the point. I just want to say I appreciate them for who they are. v Publications (Papers published, in press, or submitted, arising from the study presented in this thesis) Proche§ ~, Marshall DJ. 2001. Global distribution patterns of non-halacarid manne intertidal mites: implications for their origins in marine habitats. Journal of Biogeography 28: 47-58. Proche§ ~, Marshall DJ, Ugrasen K, Ramcharan A. 2001. Mangrove pneumatophore arthropod assemblages and seasonality patterns. Journal of the Marine Biological Association of the United Kingdom 81 : 545-552. Proche§ ~. 2001. Back to the sea: secondary marine organisms from a biogeographical perspective. Biological Journal of the Linnean Society 74: 197-203. Proche§ ~. 2001. Halacaropsis praecognita n. sp. (Acari: Halacaridae) from southern Africa. Transactions of the Royal Society ofSouth Africa, in press. Proche§ ~. 2001. New species of Copidognathinae (Acari: Halacaridae) from southern Africa. Journal ofNatural History, in press. Proche§ ~, Marshall DJ. 2001. Ecological patterns of two cohabiting evolutionary distinct animal groups: mangrove pneumatophore arthropods as a case study. Ecography, submitted. Proche§ ~, Marshall DJ. 2001. Algal growth and sediment deposition as determinants of arthropod distribution and abundance on mangrove pneumatophores. Estuarine, Coastal and Shelf Science, submitted. Proche§ ~, Marshall DJ. 2001. Diversity and biogeography of southern African marine Acari. Journal ofBiogeography , submitted. VI Contents Abstract ........ .... ...... ....... .. ... ... .. ... ...... .... ..... ....... ... ... .... ......... ................. 111 Acknowledgements .. .. .. ....... .... ... ..... ........ .. ... .............. .... ... .. .. .... ......... .... .. .v Publications .. ..... ...... .. ... ..... .... .................... ..... ...... ... ....... .... .. ........... .. .... VI Table of contents ... ..... .. ...... ... .... ... ...... ... ........ ... ... .. ····.· ... ... .. ···. ···· ····· ···· .V11 Chapter I. Introduction ....... .... ... ................ .. ............ ....... ................ ............ .1 Section I. Biogeography Chapter II. Back to the sea: secondary marine organisms from a biogeographical perspective .. .... .... ..... ..... .................. ........... ...... ....... ......... ...... ... .. .. 10 Chapter III. Global distribution patterns of non-halacarid marine intertidal mites: implications for their origins in marine habitats .......... ... ..... .. .. ........ ... .. .. ... 25 Chapter IV. Diversity and biogeography of southern African intertidal Acari ............. .... 50 Section II. Ecology Chapter V. Mangrove pneumatophore arthropod assemblages and temporal patterns .. .... .. ..79 Chapter VI. Assemblage structure of two phylogenetically distinct arthropod
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